Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia.

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.